Many of the bolded characters in the characterization above are apomorphies of subsets of streptophytes along the lineage leading to the embryophytes, not apomorphies of crown-group embryophytes per se.

All groups below are crown groups, nearly all are extant. Characters mentioned are those of the immediate common ancestor of the group, [] contains explanatory material, () features common in clade, exact status unclear.

Age. Magallón and Castillo (2009: but topology) offer estimates of ca 93.75 m.y. for the crown group, while ages in Beaulieu et al. (2013a) are ca 93 m.y. and in Magallón et al. (2015) ca 86.8 m.y. old.
Note that many estimates of stem-group ages for Bruniales - implicitly the age of this node - are based on very different topologies that that here, and/or ignore the existence of Columelliaceae as sister to Bruniaceae.

Evolution.Divergence & Distribution. Diversification at this node probably occurred in the southern hemisphere (Beaulieu et al. 2013a). Bruniales are another old, species poor but morphologically quite diverse clade that is placed along the spine of the campanulids.

Phylogeny. For the relationships of Bruniales, see the asterid II clade.

Note: Possible apomorphies are in bold. However, the actual level at which many of these features, particularly the more cryptic ones, should be assigned
is unclear. This is partly because many characters show considerable homoplasy, in addition, basic information for all too many is very incomplete, frequently coming from taxa well embedded in the clade of interest and so making the position of any putative apomorphy uncertain. Then there is the not-so-trivial issue of how ancestral states are reconstructed (see above).

Phylogeny. Within Bruniales, Bruniaceae are sister (1.0 Bayesian p.p.) to Columelliaceae in many analyses in Winkworth et al. (2008a), although not when coding chloroplast sequences were used alone, while Soltis et al. (2011) found strong support for this clade only when mitochondrial genes were removed from the analysis. Both palynology and wood anatomy had suggested a relationship between Columellia and Desfontainia (ex Loganiaceae) (Mennenga, in Leeuwenberg 1980), and this was strongly supported by rbcL and other
data (see B. Bremer et al. 1994; Backlund & Bremer 1997; Bell et al. 2001).

Chemistry, Morphology, etc. For more on the inferior/superior ovary distinction in the campanulids, see the Asterales page.

Classification. Allthough Columellia and Desfontainea at first sight look rather different, they have a number of similarities, including some possible apomorphies, in common, and the circumscription adopted here seems reasonable.

Previous Relationships. In the first seven versions of this site (pre April 2008) Columelliaceae s.l. (= [Columelliaceae + Desfontainiaceae]) were placed immediately before the ordinal characterisation of Dipsacales - it seemed to make morphological sense and there was some other evidence for this position.

Bruniaceae are ericoid shrubs with closely-set, spirally-arranged, needle-like leaves with entire margins which often have a black projection at the apex; it is restricted to the Cape region of South Africa. The often small flowers are frequently aggregated into more or less capitate inflorescences, the adaxial surfaces of the petals showing a diversity of swellings or ridges. The ovary is more or less inferior.

Evolution.Divergence & Distribution. The three main clades (tribes) had all diverged by 33.7-20.2 m.y.a., yet much of the diversification within them has occurred within the last 18-3 m.y. (Quint & Claßen-Bockhoff 2008).

Chemistry, Morphology, etc. Bruniaceae are poorly known. The plant is reported to be scented and the leaf apex has suberised cells produced by a localised
cork cambium, hence the black tip (Carlquist 1990a). The corolla "tube" is at least sometimes
formed by adnation of the filaments to adjacent free petals, and the petals are developmentally initally free (Quint & Claßen-Bockhoff 2006b). The androecium is often weakly
monosymmetric, the abaxial pair of stamens being larger than the others. Gynoecial variation is considerable; in genera like Berzelia where there is only a single carpel (really pseudomonomery?), there is only a single locule, ovule and style. There are only one or two layers of parietal cells.

Classification. Claßssen-Bockhoff et al. (2011) propose a tribal classification based on the phylogeny by Quint and Claßssen-Bockhoff (2006a). For a somewhat dated monograph, see Pillans (1947).

Previous Relationships. Bruniaceae have often been linked with the South African Grubbiaceae, here in Cornales (see Hall 1987 for some references), but the similarities between the two probably reflect the fact that they are ericoid shrubs growing in similar habitats.

Evolution.Divergence & Distribution. Another old, species-poor but morphologically quite diverse group that is restricted to South America.

Chemistry, Morphology, etc. The intervascular pits of Desfontainia are
scalariform or circular and bordered, not vestured. All the cells in even the young stem are slightly lignified. Myricetin?

The two stamens of Columellia are the adaxial pair and have latrorse, semicircular thecae attached their length to an expanded
connective; the anthers are
described as being extrorse by Backlund and Donoghue (1996). Although both Desfontainia and Viburnum have similar, smooth pollen orbicules, these are also quite widely distributed in Gentianales, at least (Vinckier & Smets 2002). Maldonado de Magnano (1986a) suggests that the ovules of Desfontainea are weakly
crassinucellate, with a single layer of nucellar cells between the megaspore and the nucellar epithelium; an endothelium is at most
poorly developed, and there is a large haustorial suspensor.

For Columellia information was taken from Hasselberg (1937), Stern et al. (1969: anatomy), Backlund and Donoghue (1996: general), and Gregory (1998: anatomy), for waxes,
see Theisen and Barthlott (1994) and Fehrenbach and Barthlott (1988: cuticle platelets
as ribbons and rodlets): Further data from: Zak & Jaramillo 3266. For Desfontainia information was taken from
Backlund and Donoghue (1996) and Hasselberg (1937): Further data from: Qin 710 and Zarucchi et al. 5195.

Previous Relationships. The relationships of Columelliaceae have long been uncertain; the family was placed in Rosales by Cronquist
(1981) and in Hydrangeales by Takhtajan (1997). Desfontainea has usually been included in the heterogeneous
Loganiaceae - and so entirely unrelated.